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VI Vaccination of Non-Domestic Carnivores Transmissible Diseases Handbook VI. Vaccination of Non-Domestic Carnivores : a Review Joost Philippa, DVM Institute of Virology, Erasmus MC, Rotterdam, The Netherlands Since the introduction of vaccinia by Jenner 200 years ago, vaccines have been the first line of defense in controlling infectious diseases in man and domestic animals. Large scale domestic dog vaccination programmes against canine distemper virus (CDV) and canine adenovirus (CAV) became possible in the late 1940’s, when egg-adapted vaccines became available on a commercial basis, followed by tissue culture adapted vaccines in the late 1950’s (Piercy 1961). Before this time these devastating diseases had to be controlled through quarantine and vigilance in capturing feral domestic animals (Dolensek et al. 1977). Historically there have always been two major types of vaccines based on the living state of the antigens. Modified live (MLV) vaccines use attenuated pathogens which reproduce in the vaccinate, thereby elliciting an immunologic response without causing disease (a “controlled” infection). The other major type of vaccine uses antigens that are non-living or inert – killed vaccine (KV). These vaccinations are preferred when safety information is not available, as they do not replicate and are therefore incapable of causing an infection. The proces of inactivation however may be damaging enough to modify immunogenicity, usually resulting in an immune response that is shorter in duration, narrower in antigenic spectrum, weaker in cell-mediated and mucosal immune responses, and possibly less effective in totally preventing viral entry (Murphy et al. 1999). Recent advances in immunology, molecular biology and biochemistry have allowed the construction of subunit vaccines based on viral or bactererial recombinants, peptides, or plasmid vectors, which may lead to safer, more efficacious vaccines that can also be used in exotic species. Vaccines cannot be absolutely guaranteed to provide protection against disease. The principal objective of vaccination is to mimic the protective immune response induced by natural infection, i.e. to elicit a high titre of neutralising antibodies of the appropriate class, IgG and/or IgA, directed against the relevant epitopes on the virion (Murphy et al. 1999). Immunity induced by vaccination or infection in domestic animals has been evaluated mainly by measuring the levels of serum antibodies. The immunologic response that provides protection against infectious agents involves a cellular and a humoral response. For certain infections (e.g. CDV, CAV, canine parvovirus (CPV), feline panleukopenia virus (FPV) or Borrelia burghdorferi ) the humoral responses, although not the only mechanism involved, tend to correlate with level of protection from clinical disease, and therefore may be a useful indicator of the immune status. Other agents (eg, Bordetella bronchiseptica , canine coronavirus (CCoV), feline enteric coronavirus (FCoV), canine parainfluenzavirus (CPIV) and Chlamydia psittaci ) all replicate and cause damage on mucosal surfaces, and might require a mucosal immune response for protection. As a consequence, serum antibody titres do not necessarily correlate with protection (Pfizer 1998). A high concentration of antibodies in an VI. Vaccination of Non-Domestic Carnivores : a Review animal implies that the animal is probably protected but also could indicate that it may not be possible to stimulate an additional immune response in that animal. Cattle vaccinated with baculovirus-expressed haemaglutinin (H) and fusion (F) proteins of Rinderpest virus (RPV) were not protected against disease caused by challenge exposure despite having detectable antibody titres (Bassiri et al. 1993). For some diseases, caused by persistent intracellular pathogens (viral diseases or intracellular bacteria) the neutralising antibodies and complement may play a less important role than the cellular immunity. Jones et al. (1993, 1997) demonstrated a lack of detectable antibody titre to CDV in ferrets and to peste des petites ruminants (PPRV) in goats after vaccination with a RPV recombinant vaccine based on fowlpox expressing the F and H genes. When these animals were challenged they survived, suggesting that protection against clinical disease may be cell mediated rather than humoral. A comparable study by Fisher et al (2003) using a DNA vaccine showed clear protection after challenge with CDV, with only limited virus neutralising antibody titres. High antibody concentrations do serve to inhibit the spread of virus between cells and thus promote host resistance (Tizard & Ni 1998). Infection with feline rhinotracheitis virus, as all herpesviruses, requires local and cell-mediated immunity, and one study suggests that there is no correlation with protective immunity and antibody titre (Johnson & Povey 1985). Later studies (eg Scott & Geissinger 1999, Dawson et al. 2001) use antibody titres as indicators of protective effect. In general it can be stated that theoretically, cell-mediated immunity is the most effective arm of the immune system in controlling, if not eliminating latent/persistent infections such as those caused by herpesviruses and retroviruses (Murphy et al. 1999). The usefulness of antibody titres to measure immunity is thus limited to only a few disease agents, and in order to obtain a more complete view of the immunologic status of an animal one therefore needs to look at the humoral and cellular responses. One should also keep in mind that an animal that has mounted an immune response after vaccination will possess memory T and B cells, which will remain for years after the antibody titre has declined. These memory cells rapidly differentiate during a subsequent infection into effector cells that can eliminate an infection before clinical signs appear, although the exact mechanism responsible for this longevity is unknown (Ahmed & Gray 1996). One can only know if the measured level of immunity is protective by challenging the vaccinated animal with the pathogen. One of the principal causes of vaccination failures in domestic dogs is maternal antibody interference. MLV vaccines differ in their ability to evade antibodies, and may sometimes be prevented from inducing an immunologic response in the vaccinate when the antibody level is high. The duration of passive immunity is directly correlated with the metabolic size of the animal. Therefore immunoglobulins will persist longer in a larger animal (Armstrong et al. 1942). The “window of vulnerability” during which the pup is vulnerable to infection with virulent virus, but unresponsive to attenuated vaccine virus (Pollock and Carmichael 1990), has been shown to range from 2-5 weeks for parvovirus infection in domestic dogs, but varies between pathogens and vaccinate species. Females close to parturition may be hyperimmunised with an inactivated parvovirus vaccine, so that high levels of maternal antibody delay the window of vulnerability until the offspring are older and better able to withstand the effects of parvovirus infection. The recent debate in veterinary medicine concerning issues related to vaccine efficacy and safety as well as duration of immunity induced by the currently available vaccines (Smith et al. 1995, Schultz et al. 1998, Kruth & Ellis 1998, Tizard & Ni 1998, McCaw et al. 1998, Gumley et al. 1999, Hustead et al. 1999, Twark et al. 2000) has resulted in the need for more objective and scientific data and an increase in research in domestic animals. In non-domestic animals however, there have been only few controlled studies of vaccination (Heerden et al. 1980, Halbrooks et al. 1981, Behlert et al. 1981, Barker et al. 1983, Montali et al. 1983, Green et al. 1984, Bush et al. 1985, Hoover et al. 1985, Paul- Murphy et al. 1985, Briggs et al. 1986, Tham et al. 1987, Follmann et al. 1988, Spencer et al. 1991, 1992, Goodrich et al. 1994, Harrenstien et al. 1995, Schubert et al. 1995, Williams et al. 1996, Henke et al. 1997, Kadoi et al. 1998, Bingham et al. 1999, Harthorn et al. 1999, Pare et al. 1999, Wimsatt et al. 1999, 2003, Maack et al. 2000, Blasco et al. 2001, Federhoff VI. Vaccination of Non-Domestic Carnivores : a Review et al. 2001, Lambot et al. 2001, Maia et al. 2001, van Heerden et al. 1998, 2002). These are usually restricted to measuring the (humoral) immune response and extrapolating the data from those known in domestic animals, as subjecting (endangered) zoo animals to challenge infections is generally not an option. Challenge studies in dogs have shown a range of reported protective titres – these may differ due to the variety of techniques and standards used. In humans there is a general standardisation of assay methods to measure antibody titres. Non-standardisation of serologic tests makes comparisons between laboratories of questionable use (Luff et al. 1987): Reported protective titres in domestic dogs : virus protective Reference titre CAV ≥30 Cole et al. 1998 CDV >2 Gillespie et al. 1965; Ackerman & Siebel 1974. ≥20 Gillespie et al. 1958, 1972;Gorham et al. 1966; Prydie 1966; Krakowka et al. 1978; Cooper et al. 1991. ≥24 Jones et al. 1997. ≥30 Gillespie 1966. >50 Olson et al. 1988. ≥96 Mc Caw et al. 1998. ≥100 Appel 1969; Krakowka et al. 1975, Montali et al. 1983; Carmichael et al. 1999. CPV ≥80 Olson et al. 1988; Carmichael et al. 1983, 1994, 1997; Appel et al. 1979; Pollock and Carmichael 1982; Meunier et al. 1985. RV ≥20 Bunn et al. 1984 Twark et al. (2000) report that CDV titres above 5 are indicative of an adequate antibody response in domestic dogs, although level of protection is unknown, and recommend CDV revaccination when titres are below 32. Carmichael et al. (1983) demonstrated that dogs with a CPV titre below 100 were not protected. Titres of 200-800 were protective in some dogs, titres above 1600 appeared to be protective in all dogs. There is still no general consensus on how often domestic animals need to be revaccinated - for most vaccines there is little information on the duration of immunity. It is, however, recognised that protective immunity to CDV following MLV vaccination is of long duration, perhaps even lifelong. For other viruses or components of combination vaccines this duration may not be of such long duration.
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